Biomics

After nearly half a century of dominance by the Oparin–Haldane coacervate hypothesis, which posited the primacy of peptides in the origin of life, researchers gradually shifted their focus to another type of biopolymer — namely, RNA molecules, which led to the assumption that the planet initially had an RNA world. Despite the fact that RNA is capable of exhibiting catalytic activity as ribozymes and storing certain (hereditary) information, even chemical pre‑Darwinian evolution would not have made much progress without interaction with peptides. It can be considered established that amino acids formed from atmospheric gases under the influence of appropriate external factors (lightning, ultraviolet radiation, thermal energy), and through condensation they formed peptides. Another type of polymeric molecules in the form of RNA arose through the synthesis of nitrogenous bases from hydrocyanic acid, which combined with a carbohydrate component in the form of ribose. Ribose originated from formaldehyde during the formose reaction (and, to increase stability, formed a complex with borates); after that, the resulting nucleosides reacted with phosphates, ultimately leading to the formation of nucleotides, which, in turn, underwent condensation and created RNA molecules. Some of these reactions took place in the gas, liquid, and/or solid phases. The next stage was the formation of ribonucleoproteins (RNPs), which are complexes of peptides and RNA. The evolution of RNP led to the emergence of a peptidyltransferase center capable of catalyzing the formation of peptide bonds, after which the genetic code arose—but how this happened remains the most serious mystery. Subsequent compartmentalization and the formation of ribosomes served as an important impetus for the emergence of protocells and the first primitive structures—not yet organisms, but certain formations that came to be known as “progenotes.” At this stage, DNA appeared, initially carrying uracils, which were later replaced by thymines. It was the formation of DNA molecules that became a landmark event, taking the storage and transmission of hereditary information to a new level, while leaving RNA and proteins to perform catalytic and structural functions — which can now be represented as the DNA/RNA/protein triad. A hypothetical scenarios of abiotic chemical evolution was proposed, followed by Darwinian evolution of biopolymers, which led to the emergence of modern life. Understanding how life emerged on Earth is one of the greatest problems in natural science and has a fundamental character. However, with the emergence of a new field in biology – “synthetic biology” – the quest to understand the origin of life has ceased to be of idle interest and has gone beyond the scope of purely fundamental research, yet it has not yet moved into the practical realm.

Origin of Life, amino acids, peptides, proteins, ribose, nitrogenous bases, nucleoside, nucleotide, RNA, deoxyribose, DNA

1. Abramov O, Medvegy A, Kremer B, Mojzsis SJ. A Hadean timeline for the emergence of the RNA World. Nat Commun. 2026 Sep 22;17(1):9790. doi: 10.1038/s41467-026-76978-3
2. Alekseev Ya.I., Garafutdinov R.R., Chemeris D.A., Sharipov B.T., Vorob`ev A.A., Zhevora S.V., Kuzubov A.V., Chemeris A.V. Chemical synthesis of oligonucleotides: stages of a 70-year journey. Biomics. 2026. 18(1). 178-214. doi: 10.31301/2221-6197.bmcs.2026-14 (In Russian)
3. Anosova I, Kowal EA, Sisco NJ et al. Structural Insights into Conformation Differences between DNA/TNA and RNA/TNA Chimeric Duplexes. Chembiochem. 2016. 17(18). 1705-1708. doi: 10.1002/cbic.201600349
4. Attwater J, Tagami S, Kimoto M et al. Chemical fidelity of an RNA polymerase ribozyme. Sci. 2013; 4 (7): 2804–2814. doi: 10.1039/c3sc50574j
5. Attwater J, Wochner A, Pinheiro VB et al. Ice as a protocellular medium for RNA replication. Nat Commun. 2010. 1. 76. doi: 10.1038/ncomms1076
6. Baaske P, Weinert FM, Duhr S et al. Extreme accumulation of nucleotides in simulated hydrothermal pore systems. Proc Natl Acad Sci USA. 2007. 104(22). 9346-9351. doi: 10.1073/pnas.0609592104
7. Bada JL. Volcanic Island lightning prebiotic chemistry and the origin of life in the early Hadean eon. Nat Commun. 2023. 14(1). 2011. doi: 10.1038/s41467-023-37894-y
8. Bada JL, Bigham C, Miller SL et al. Impact melting of frozen oceans on the early Earth: implications for the origin of life. Proc Natl Acad Sci USA. 1994. 91(4). 1248-1250. doi: 10.1073/pnas.91.4.1248
9. Bánfalvi G. DNA Empire. In: Apoptotic Chromatin Changes. Springer, Dordrecht. 2009. 1-30. doi: 10.1007/978-1-4020-9561-0_1
10. Banfalvi G. Ribose Selected as Precursor to Life. DNA Cell Biol. 2020. 39(2). 177-186. doi: 10.1089/dna.2019.4943
11. Banfalvi G. Prebiotic Pathway from Ribose to RNA Formation. Int J Mol Sci. 2021. 22(8). 3857. doi: 10.3390/ijms22083857
12. Bekker A, Holland HD, Wang PL et al. Dating the rise of atmospheric oxygen. Nature. 2004. 427(6970). 117-120. doi: 10.1038/nature02260
13. Bernal JD. The Physical Basis of Life. Proceedings of the Physical Society. Section A. 62(9). 537-558.
14. Bernhardt HS. The RNA world hypothesis: the worst theory of the early evolution of life (except for all the others)(a). Biol Direct. 2012. 7. 23. doi: 10.1186/1745-6150-7-23
15. Bernhardt HS, Sandwick RK. Purine biosynthetic intermediate-containing ribose-phosphate polymers as evolutionary precursors to RNA. J Mol Evol. 2014. 79(3-4). 91-104. doi: 10.1007/s00239-014-9640-1
16. Bhowmik S, Krishnamurthy R. The role of sugar-backbone heterogeneity and chimeras in the simultaneous emergence of RNA and DNA. Nat Chem. 2019. 11(11). 1009-1018. doi: 10.1038/s41557-019-0322-x
17. Bielski R, Tencer M. Possible Enantioseparation of Racemic Ribose on Chiral Surface Formed by Adsorption of Nucleobases. Life (Basel). 15(8). 1160. doi: 10.3390/life15081160
18. Bonner WA. The origin and amplification of biomolecular chirality. Orig Life Evol Biosph. 1991. 21(2). 59-111. doi: 10.1007/BF01809580
19. Brachet J. Les acides nucléiques et l’origène des protéines. In: The origin of life on Earth (Academy of Sciences of the U.S.S.R: Oparin AI, Pasynskii AG, Braunshtein AE, Pavlovskaya TE (eds), English-French-German edition, Clark F, Synge RLM (eds)). MacMillan, New York, 361–367. (In French)
20. Bregestowski PD. “RNA World", a highly improbable scenario of the life origin and early ecolution on Earth J Evol Biocgem Physiol. 2015. 51(1). 64-74. doi: 10.1134/S0022093015010111
21. Breslow R. On the mechanism of the formose reaction. Tetrahedron Letters. 1959. 1(21). 22-26. doi: 10.1016/S0040-4039(01)99487-0.
22. Butlerow A. Bildung einer zuckerartigen Substanz durch Synthese. Justus Liebigs Annalen der Chemie. 1861. 120(3). 295-298. doi: 10.1002/jlac.18611200308
23. Caetano-Anollés G. The proteomic origin of the genetic code. Expert Rev Proteomics. 2026. 23(3). 79-98. doi: 10.1080/14789450.2026.2646677
24. Chaput JC, Herdewijn P. What Is XNA? Chem. Int. Ed. 2019. 58(34). 11570-11572. doi: 10.1002/anie.201905999
25. Chaput JC, Szostak JW. TNA synthesis by DNA polymerases. J Am Chem Soc. 2003. 125(31). 9274-9275. doi: 10.1021/ja035917n
26. Chargaff E. Nucleic acids as carriers of biological information. In: The Origin of Life on Earth: Collection of Reports at the International Meeting, August 1957, Moscow. : Publishing House of the Academy of Sciences of the USSR. 1957. 209-213. 192-197. (In Russian)
27. Cleaves HJ, Chalmers JH, Lazcano A et al. A reassessment of prebiotic organic synthesis in neutral planetary atmospheres. Orig Life Evol Biosph. 2008. 38(2). 105-115. doi: 10.1007/s11084-007-9120-3
28. Costanzo G, Saladino R, Botta G et al. Generation of RNA molecules by a base-catalysed click-like reaction. Chembiochem. 2012. 13(7). 999-1008. doi: 10.1002/cbic.201200068
29. Costanzo G, Saladino R, Crestini C et al. Formamide as the main building block in the origin of nucleic acids. BMC Evol Biol. 2007. 7(2). S1. doi: 10.1186/1471-2148-7-S2-S1
30. Crick FH. The origin of the genetic code. J Mol Biol. 1968. 38(3). 367-379. doi: 10.1016/0022-2836(68)90392-6
31. Crick F. Central dogma of molecular biology. Nature. 1970. 227(5258). 561-563. doi: 10.1038/227561a0
32. Dalrymple GB. The age of the Earth in the twentieth century: a problem (mostly) solved. Geological Society, London, Special Publications. 2001. 190. 205 – 221. doi: 10.1144/GSL.SP.2001.190.01.14
33. de Farias TS, Rêgo GT, José MV. Peptidyl Transferase Center and the Emergence of the Translation System. Life (Basel). 2017. 7(2). 21. doi: 10.3390/life7020021
34. Di Giulio M. On the RNA world: evidence in favor of an early ribonucleopeptide world. J Mol Evol. 1997. 45(6). 571-578. doi: 10.1007/pl00006261
35. Eschenmoser A. Chemical etiology of nucleic acid structure. Science. 1999. 284(5423). 2118-2124. doi: 10.1126/science.284.5423.2118
36. Eigen M. Selforganization of matter and the evolution of biological macromolecules. Naturwissenschaften. 1971. 58(10). 465-523. doi: 10.1007/BF00623322
37. Eigen M, Schuster P. The hypercycle. A principle of natural self-organization. Part A: Emergence of the hypercycle. Naturwissenschaften. 1977. 64(11). 541-565. doi: 10.1007/BF00450633
38. Ekland EH, Bartel DP. RNA-catalysed RNA polymerization using nucleoside triphosphates. Nature. 1996. 382(6589). 373-376. doi: 10.1038/382373a0
39. Ertem G, Ferris JP; New Collective Author. Synthesis of RNA oligomers on heterogeneous templates. Nature. 1996 Jan 18;379(6562):238-40. doi: 10.1038/379238a0
40. Farias ST, Prosdocimi F. RNP-world: The ultimate essence of life is a ribonucleoprotein process. Genet Mol Biol. 2022. 45(3) e20220127. doi: 10.1590/1678-4685-GMB-2022-0127
41. Farias ST, Rêgo TG, José MV. Origin and evolution of the Peptidyl Transferase Center from proto-tRNAs. FEBS Open Bio. 2014. 4. 175-178. doi: 10.1016/j.fob.2014.01.010
42. Ferris JP, Ertem G, Kamaluddin et al. Mineral catalysis of the formation of the phosphodiester bond in aqueous solution: the possible role of montmorillonite clays. Adv Space Res. 1989. 9(6). 67-75. doi: 10.1016/0273-1177(89)90210-x
43. Ferris JP, Kamaluddin. Oligomerization reactions of deoxyribonucleotides on montmorillonite clay: the effect of mononucleotide structure on phosphodiester bond formation. Orig Life Evol Biosph. 1989. 19. 609-619. doi: 10.1007/BF01808121
44. Ferris JP, Sanchez RA, Orgel LE. Studies in prebiotic synthesis. 3. Synthesis of pyrimidines from cyanoacetylene and cyanate. J Mol Biol. 1968. 33(3). 693-704. doi: 10.1016/0022-2836(68)90314-8
45. Forterre P. Genomics and early cellular evolution. The origin of the DNA world. C R Acad Sci III. 2001. 324(12). 1067-1076. doi: 10.1016/s0764-4469(01)01403-2
46. Forterre P. The two ages of the RNA world, and the transition to the DNA world: a story of viruses and cells. Biochimie. 2005. 87(9-10). 793-803. doi: 10.1016/j.biochi.2005.03.015
47. Forterre P. Why are there so many diverse replication machineries? J Mol Biol. 2013. 425(23). 4714-4726. doi: 10.1016/j.jmb.2013.09.032
48. Forterre P. The Last Universal Common Ancestor of Ribosome-Encoding Organisms: Portrait of LUCA. J Mol Evol. 2024. 92(5). 550-583. doi: 10.1007/s00239-024-10186-9
49. Furukawa Y, Horiuchi M, Kakegawa T. Selective stabilization of ribose by borate. Orig Life Evol Biosph. 2013. 43(4-5). 353-361. doi: 10.1007/s11084-013-9350-5
50. Gamow G. Possible Relation between Deoxyribonucleic Acid and Protein Structures. Nature. 1954. 173. 318. doi: 10.1038/173318a0
51. Gianni E, Kwok SLY, Wan CJK et al. A small polymerase ribozyme that can synthesize itself and its complementary strand. Science. 2026. 391(6789). 1022-1028. doi: 10.1126/science.adt2760
52. Gilbert W. Origin of life: The RNA world. Nature. 1986. 319(6055). 618. doi: 10.1038/319618a0
53. Griffith RW. A specific scenario for the origin of life and the genetic code based on peptide/oligonucleotide interdependence. Orig Life Evol Biosph. 2009. 39(6). 517-531. doi: 10.1007/s11084-009-9169-2
54. Hanczyc MM, Mansy SS, Szostak JW. Mineral surface directed membrane assembly. Orig Life Evol Biosph. 2007. 37(1). 67-82. doi: 10.1007/s11084-006-9018-5
55. Herdewijn P. TNA as a Potential Alternative to Natural Nucleic Acids. Angew Chem Int Ed Engl. 2001. 40(12). 2249-2251. doi: 10.1002/1521-3773(20010618)40:12<2249::aid-anie2249>3.0.co;2-i
56. Hirakawa Y, Kakegawa T, Furukawa Y. Borate-guided ribose phosphorylation for prebiotic nucleotide synthesis. Sci Rep. 2022. 12(1). 11828. doi: 10.1038/s41598-022-15753-y
57. Huang W, Ferris JP. One-step, regioselective synthesis of up to 50-mers of RNA oligomers by montmorillonite catalysis. J Am Chem Soc. 2006. 128(27). 8914-8919. doi: 10.1021/ja061782k
58. Ikehara K. [GADV]-protein world hypothesis on the origin of life. Orig Life Evol Biosph. 2014. 44(4). 299-302. doi: 10.1007/s11084-014-9383-4
59. Joshi PC, Aldersley MF, Ferris JP. Progress in demonstrating total homochiral selection in montmorillonite-catalyzed RNA synthesis. Biochem Biophys Res Commun. 2011. 413(4). 594-598. doi: 10.1016/j.bbrc.2011.09.008
60. Joshi PC, Pitsch S, Ferris JP. Selectivity of montmorillonite catalyzed prebiotic reactions of D, L-nucleotides. Orig Life Evol Biosph. 2007. 37(1). 3-26. doi: 10.1007/s11084-006-9013-x
61. Joyce GF, Schwartz AW, Miller SL, Orgel LE. The case for an ancestral genetic system involving simple analogues of the nucleotides. Proc Natl Acad Sci USA. 1987. 84(13). 4398-4402. doi: 10.1073/pnas.84.13.4398
62. Koonin EV. Frozen Accident Pushing 50: Stereochemistry, Expansion, and Chance in the Evolution of the Genetic Code. Life (Basel). 7(2). 22. doi: 10.3390/life7020022
63. Koonin EV, Novozhilov AS. Origin and Evolution of the Universal Genetic Code. Annu Rev Genet. 2017. 51. 45-62. doi: 10.1146/annurev-genet-120116-024713
64. Kondratyeva LG, Dyachkova MS, Galchenko AV. The Origin of Genetic Code and Translation in the Framework of Current Concepts on the Origin of Life. Biochemistry (Mosc). 2022 Feb;87(2):150-169. doi: 10.1134/S0006297922020079
65. Kruse FM, Teichert JS, Trapp O. Prebiotic Nucleoside Synthesis: The Selectivity of Simplicity. Chemistry. 2020. 26(65). 14776-14790. doi: 10.1002/chem.202001513
66. Kun Á, Radványi Á. The evolution of the genetic code: Impasses and challenges. Biosystems. 2018. 164. 217-225. doi: 10.1016/j.biosystems.2017.10.006
67. Larralde R, Robertson MP, Miller SL. Rates of decomposition of ribose and other sugars: implications for chemical evolution. Proc Natl Acad Sci USA. 92(18). 8158-8160. doi: 10.1073/pnas.92.18.8158
68. Lawrence MS, Bartel DP. New ligase-derived RNA polymerase ribozymes. RNA. 2005. 11(8). 1173-1180. doi: 10.1261/rna.2110905
69. Le Vay K, Mutschler H. The difficult case of an RNA-only origin of life. Emerg Top Life Sci. 2019. 3(5). 469-475. doi: 10.1042/ETLS20190024
70. Lesk AM. Why does DNA contain thymine and RNA uracil? J Theor Biol. 1969. 22(3). 537-540. doi: 10.1016/0022-5193(69)90021-6
71. Lessor RA, Leonard NJ. Synthesis of 2'-deoxynucleosides by deoxygenation of ribonucleosides. Journal of Organic Chemistry. 1981. 46(21). 4300–4301. doi: 10.1021/jo00334a043
72. Loew O. Einige Bemerkungen über Formose. Dtsch. Chem. Ges. 1887. 20. 141–144.
73. Mallik S, Kundu S. The Lipid-RNA World. 2012. 1211.0413. DOI: 10.48550/arXiv.1211.0413
74. Martin W, Baross J, Kelley D, Russell MJ. Hydrothermal vents and the origin of life. Nat Rev Microbiol. 2008. 6(11). 805-814. doi: 10.1038/nrmicro1991
75. Mast CB, Schink S, Gerland U, Braun D. Escalation of polymerization in a thermal gradient. Proc Natl Acad Sci USA. 2013. 110(20). 8030-8035. doi: 10.1073/pnas.1303222110
76. Matsuo M, Kurihara K. Proliferating coacervate droplets as the missing link between chemistry and biology in the origins of life. Nat Commun. 2021. 12(1). 5487. doi: 10.1038/s41467-021-25530-6
77. Miller SL. A production of amino acids under possible primitive earth conditions. Science. 1953. 117(3046). 528-529. doi: 10.1126/science.117.3046.528
78. Miller SL, Schopf JW, Lazcano A. Oparin's "Origin of Life": sixty years later. J Mol Evol. 1997. 44(4). 351-353.
79. Miller SL, Urey HC. Organic compound synthesis on the primitive earth. Science. 1959. 130(3370). 245-251. doi: 10.1126/science.130.3370.245
80. Miyakawa S, Ferris JP. Sequence- and regioselectivity in the montmorillonite-catalyzed synthesis of RNA. J Am Chem Soc. 2003. 125(27). 8202-8208. doi: 10.1021/ja034328e
81. Mojzsis SJ, Arrhenius G, McKeegan KD et al. Evidence for life on Earth before 3,800 million years ago. Nature. 1996. 384(6604). 55-59. doi: 10.1038/384055a0
82. Mojzsis SJ, Harrison TM, Pidgeon RT. Oxygen-isotope evidence from ancient zircons for liquid water at the Earth's surface 4,300 Myr ago. Nature. 2001. 409(6817). 178-181. doi: 10.1038/35051557
83. Moldavanov A. Physical factors in origin of genetic code. Biosystems. 2026. 262. 105712. doi: 10.1016/j.biosystems.2026.105712
84. Moldogazieva NT, Terentiev AA, Mokhosoev IM, Astakhov DV. Redox chemistry of early Earth and the origin of life. Commun Chem. 2026. 9(1). 143. doi: 10.1038/s42004-026-01969-w
85. Mulkidjanian AY, Cherepanov DA, Galperin MY. Survival of the fittest before the beginning of life: selection of the first oligonucleotide-like polymers by UV light. BMC Evol Biol. 2003. 3. 12. doi: 10.1186/1471-2148-3-12
86. Mulkidjanian AY, Dibrova DV, Bychkov AY. Origin of the RNA World in Cold Hadean Geothermal Fields Enriched in Zinc and Potassium: Abiogenesis as a Positive Fallout from the Moon-Forming Impact? Life (Basel). 15(3). 399. doi: 10.3390/life15030399
87. Nagy KK, Skurnik M, Vértessy BG. Viruses with U-DNA: New Avenues for Biotechnology. Viruses. 2021. 13(5). 875. doi: 10.3390/v13050875
88. Nielsen PE. Peptide nucleic acids and the origin of life. Chem Biodivers. 2007. 4(9). 1996-2002. doi: 10.1002/cbdv.200790166
89. Niether D, Afanasenkau D, Dhont JK, Wiegand S. Accumulation of formamide in hydrothermal pores to form prebiotic nucleobases. Proc Natl Acad Sci USA. 2016. 113(16). 4272-4277. doi: 10.1073/pnas.1600275113
90. Orgel LE. Evolution of the genetic apparatus. J Mol Biol. 1968. 38(3). 381-393. doi: 10.1016/0022-2836(68)90393-8
91. Orgel L. Origin of life. A simpler nucleic acid. Science. 2000. 290(5495). 1306-1307. doi: 10.1126/science.290.5495.1306
92. Oró J. Synthesis of adenine from ammonium cyanide. Biochemical and Biophysical Research Communications. 1960. 2(6). 407-412. doi: 10.1016/0006-291X(60)90138-8.
93. Oró J. Mechanism of synthesis of adenine from hydrogen cyanide under possible primitive earth conditions. Nature. 1961. 191. 1193-1194. doi: 10.1038/1911193a0
94. Oró, J.; Cox, A. C. Non-Enzymic Synthesis of 2-Deoxyribose. Proc. 1962. 21. A80.
95. Papastavrou N, Horning DP, Joyce GF. RNA-catalyzed evolution of catalytic RNA. Proc Natl Acad Sci USA. 2024. 121(11). e2321592121. doi: 10.1073/pnas.2321592121
96. Poole AM, Horinouchi N, Catchpole RJ et al. The case for an early biological origin of DNA. J Mol Evol. 79(5-6). 204-212. doi: 10.1007/s00239-014-9656-6
97. Poole A, Penny D, Sjöberg B. Methyl-RNA: an evolutionary bridge between RNA and DNA? Chem Biol. 2000. 7(12). R207-216. doi: 10.1016/s1074-5521(00)00042-9
98. Poole A, Penny D, Sjöberg BM. Confounded cytosine! Tinkering and the evolution of DNA. Nat Rev Mol Cell Biol. 2001. 2(2):147-51. doi: 10.1038/35052091
99. Powner MW, Zheng S-L, Szostak J.W. Multicomponent Assembly of Proposed DNA Precursors in Water. Journal American Chemical Society. 2012. 134(33). 13889-13895. doi: 10.1021/ja306176n
100. Prieur BE. Étude de l’Activité Prébiotique Potentielle de l’Acide Borique. R. Acad. Sci., Ser. IIc: Chim. 2001. 4. 667–670. doi: 10.1016/S1387-1609(01)01266-X (In French)
101. Prosdocimi F, Farias ST. Coacervates meet the RNP-world: liquid-liquid phase separation and the emergence of biological compartmentalization. Biosystems. 2025 Jun;252:105480. doi: 10.1016/j.biosystems.2025.105480
102. Prosdocimi F, Jheeta S, Torres de Farias S. Conceptual challenges for the emergence of the biological system: Cell theory and self-replication. Med Hypotheses. 2018 Oct;119:79-83. doi: 10.1016/j.mehy.2018.07.029
103. Prosdocimi F, José MV, de Farias ST. The Theory of Chemical Symbiosis: A Margulian View for the Emergence of Biological Systems (Origin of Life). Acta Biotheor. 2021 Mar;69(1):67-78. doi: 10.1007/s10441-020-09388-7
104. Racker E. Enzymatic synthesis of deoxypentose phosphate. Nature. 1951. 167(4245). 408-409. doi: 10.1038/167408b0
105. Reichard P. The evolution of ribonucleotide reduction. Trends Biochem Sci. 1997. 22(3). 81-85. doi: 10.1016/s0968-0004(97)01003-7
106. Ricardo A, Carrigan MA, Olcott AN, Benner SA. Borate minerals stabilize ribose. Science. 2004. 303(5655). 196. doi: 10.1126/science.1092464
107. Rich A. On the problems of evolution and biochemical information transfer. M. Kasha, B. Puhlman (Eds.), Horizons in Biochemistry. Academic Press. 1962. 103-126.
108. Ritson DJ, Sutherland JD. Conversion of biosynthetic precursors of RNA to those of DNA by photoredox chemistry. J Mol Evol. 2014. 78(5). 245-250. doi: 10.1007/s00239-014-9617-0
109. Robertson MP, Miller SL. An efficient prebiotic synthesis of cytosine and uracil. Nature. 1995. 375(6534). 772-774. doi: 10.1038/375772a0
110. Robins MJ, Wilson JS, Hansske F. Nucleic acid related compounds. 42. A general procedure for the efficient deoxygenation of secondary alcohols. Regiospecific and stereoselective conversion of ribonucleosides to 2'-deoxynucleosides. Journal American Chemical Society. 1983. 105(12). 4059-4065. doi: 10.1021/ja00350a052
111. Root-Bernstein M, Root-Bernstein R. The ribosome as a missing link in the evolution of life. J Theor Biol. 2015. 367. 130-158. doi: 10.1016/j.jtbi.2014.11.025
112. Sacerdote MG, Szostak JW. Semipermeable lipid bilayers exhibit diastereoselectivity favoring ribose. Proc Natl Acad Sci USA. 2005. 102(17). 6004-6008. doi: 10.1073/pnas.0408440102
113. Samanta B, Joyce GF. A reverse transcriptase ribozyme. Elife. 2017. 6. e31153. doi: 10.7554/eLife.31153
114. Sanchez R, Ferris J, Orgel LE. Conditions for purine synthesis: did prebiotic synthesis occur at low temperatures? Science. 1966. 153(3731). 72-73. doi: 10.1126/science.153.3731.72
115. Schopf JW. The paleobiological record of photosynthesis. Photosynth Res. 2011. 107(1). 87-101. doi: 10.1007/s11120-010-9577-1
116. Schopf JW. Pioneers of Origin of Life Studies-Darwin, Oparin, Haldane, Miller, Oró-And the Oldest Known Records of Life. Life (Basel). 14(10). 1345. doi: 10.3390/life14101345
117. Schwartz AW, Joosten H, Voet AB. Prebiotic adenine synthesis via HCN oligomerization in ice. Biosystems. 1982. 15(3). 191-193. doi: 10.1016/0303-2647(82)90003-x
118. Shapiro R. Prebiotic ribose synthesis: a critical analysis. Orig Life Evol Biosph. 1988. 18(1-2). 71-85. doi: 10.1007/BF01808782
119. Skoblikow NE, Zimin AA. Mineral Grains, Dimples, and Hot Volcanic Organic Streams: Dynamic Geological Backstage of Macromolecular Evolution. Journal of Molecular Evolution. 2018. 86(3). 172-183. 1432-1432. doi: 10.1007/s00239-018-9839-7
120. Steer AM, Bia N, Smith DK, Clarke PA. Prebiotic synthesis of 2-deoxy-d-ribose from interstellar building blocks promoted by amino esters or amino nitriles. Chem Commun (Camb). 53(75). 10362-10365. doi: 10.1039/c7cc06083a
121. Subramanian H, Brown J, Gatenby R. Prebiotic competition and evolution in self-replicating polynucleotides can explain the properties of DNA/RNA in modern living systems. BMC Evol Biol. 2020. 20(1). 75. doi: 10.1186/s12862-020-01641-4
122. Tamura K. Ribosome evolution: emergence of peptide synthesis machinery. J Biosci. 2011. 36(5). 921-928. doi: 10.1007/s12038-011-9158-2
123. Trixler F. Origin of Nucleic Acids. In: Prebiotic Chemistry and the Origin of Life. 117-137. doi: 10.1007/978-3-030-81039-9_5
124. van der Gulik PT, Speijer D. How amino acids and peptides shaped the RNA world. Life (Basel). 5(1). 230-46. doi: 10.3390/life5010230
125. Vlassov AV, Kazakov SA, Johnston BH, Landweber LF. The RNA world on ice: a new scenario for the emergence of RNA information. J Mol Evol. 2005. 61(2). 264-273. doi: 10.1007/s00239-004-0362-7
126. Wang J, Yu H. Threose nucleic acid as a primitive genetic polymer and a contemporary molecular tool. Bioorganic Chemistry. 2024. 143. 107049. doi: 10.1016/j.bioorg.2023.107049
127. Wetherill GW. Formation of the Earth. Annual Review of Earth and Planetary Sciences. 1990. 18. 205-256.
128. Wieczorek R, Dörr M, Chotera A et al. Formation of RNA phosphodiester bond by histidine-containing dipeptides. Chembiochem. 2013. 14(2). 217-223. doi: 10.1002/cbic.201200643
129. Woese CR. On the evolution of the genetic code. Proc Natl Acad Sci USA. 1965. 54(6). 1546-1552. doi: 10.1073/pnas.54.6.1546
130. Woese CR, Kandler O, Wheelis ML. Towards a natural system of organisms: proposal for the domains Archaea, Bacteria, and Eucarya. Proc Natl Acad Sci USA. 87(12). 4576-4579. doi: 10.1073/pnas.87.12.4576
131. Yadav M, Kumar R, Krishnamurthy R. Chemistry of Abiotic Nucleotide Synthesis. Chem Rev. 2020. 120(11). 4766-4805. doi: 10.1021/acs.chemrev.9b00546
132. Yang YW, Zhang S, McCullum EO, Chaput JC. Experimental evidence that GNA and TNA were not sequential polymers in the prebiotic evolution of RNA. J Mol Evol. 2007. 65(3). 289-295. doi: 10.1007/s00239-007-9017-9
133. Yarus M. Evolution of the Standard Genetic Code. J Mol Evol. 2021. 89(1-2). 19-44. doi: 10.1007/s00239-020-09983-9
134. Yu H, Zhang S, Chaput JC. Darwinian evolution of an alternative genetic system provides support for TNA as an RNA progenitor. Nat Chem. 2012. 4(3). 183-187. doi: 10.1038/nchem.1241
135. Zahnle K, Schaefer L, Fegley B. Earth's earliest atmospheres. Cold Spring Harb Perspect Biol. 2010. 2(10). a004895. doi: 10.1101/cshperspect.a004895
136. Zhao Z-R, Wang X. A plausible prebiotic selection of ribose for RNA - formation, dynamic isolation, and nucleotide synthesis based on metal-doped-clays. Chem. 2021. 7(12). 3292-3308. doi: 10.1016/j.chempr.2021.09.002